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RIUNNE FCA AR Viadopico-Vega Systematics and Biodiversity ISSN: 1477-2000 (Print) 1478-0933 (Online) Journal homepage: http://www.tandfonline.com/loi/tsab20 Systematic consideration of floral microcharacters of the South American genus Chrysolaena (Vernonieae, Asteraceae) Gisela M. Via do pico, Alvaro J. Vega & Massimiliano Dematteis To cite this article: Gisela M. Via do pico, Alvaro J. Vega & Massimiliano Dematteis (2016) Systematic consideration of floral microcharacters of the South American genus Chrysolaena (Vernonieae, Asteraceae), Systematics and Biodiversity, 14:2, 224-243, DOI: 10.1080/14772000.2015.1134699 To link to this article: http://dx.doi.org/10.1080/14772000.2015.1134699 View supplementary material Published online: 30 Jan 2016. Submit your article to this journal View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tsab20 Download by: [181.1.176.133] Date: 30 January 2016, At: 07:22 Systematics and Biodiversity (2016), 14(2): 224À243 Systematic consideration of floral microcharacters of the South American genus Chrysolaena (Vernonieae, Asteraceae) GISELA M. VIA DO PICO, ALVARO J. VEGA & MASSIMILIANO DEMATTEIS Instituto de Botanica del Nordeste (UNNE-CONICET), Casilla de Correo 209, CP 3400 Corrientes, Argentina (Received 11 August 2015; accepted 3 November 2015) Floral microcharacters of the genus Chrysolaena H. Rob. (Vernonieae, Asteraceae) were analysed in detail for the first time in order to evaluate the taxonomic position of conflictive species in the group. The results were also compared with studies carried out in species of related genera. In addition to distinctive microcharacters previously studied in some species of the genus, other characters such as trichome types of the corolla, style, anthers and cypselae have been analysed for the first time. The presence of glandular apical appendage and cypselae are common characteristics among species Chrysolaena.In addition to these, this study shows that presence of glands on the style and corollas is another consistent characteristic in the genus. However, the absence of basal stylar node would not be a diagnostic character since this varies widely among species. The results indicate that most of the microcharacters of Chrysolaena analysed are quite consistent in the genus, but they are no more consistent than the pollen morphology (type ’C’) and chromosome base number (x D 10). Until now, these last two features would be most useful for separating Chrysolaena from the related genera Lessingianthus and Lepidaploa. At species level, the results show that related species can be distinguished by the different combinations of floral microcharacters. The value of microcharacters could be increased if they are combined with other morphological, cytological, and palynological data. Keywords: anthers, crystals, cypselae, idioblasts, Lepidaploinae, style, taxonomy, trichomes Introduction With the emergence of ’The new synantherology’ (King & Robinson, 1970), dozens of new genera have The Asteraceae comprise between 1600À2000 genera been split out of several traditional genera of Asteraceae and 24,000À30,000 species divided into 13 subfamilies on the basis of micromorphological characters. In subse- and43tribes(Paneroetal.,2014). The family has pre- quent years, several authors have used a cluster of micro- sented continuing taxonomic problems as a result of its morphological characters in taxonomic studies in large size and its recurring combinations of superficial char- groups as Senecioneae, Astereae, Vernonieae, Eupator- acters (Robinson, 1999). The tribe Vernonieae is a clear ieae, Inuleae (Dematteis, 2009, Faust & Jones, 1973; Isa- Downloaded by [181.1.176.133] at 07:22 30 January 2016 example of one of the most complex and less elucidated wumi, 1999; Isawumi, El-Ghazaly, & Nordenstam, 1996; groups. It is one of the largest within the Asteraceae Liu & Yang, 2011; Narayana, 1979; Pruski, 2012; Riva, with approximately 1500À1600 species distributed in Pozner, & Freire, 2009; Robinson, 1988a, 1988b, 1992, America, Asia, and Africa (Keeley, Forsman, & Chan, 1999). Some uses of microcharacters are, for example, 2007; Keeley & Robinson, 2009). Good taxonomic species identification from small fragments of the plant, understanding of Vernonieae has been difficult to or the relocation or change of taxonomic position of taxa achieve, especially among certain New World taxa (Robinson, 2009). (Ekman, 1914; Gleason, 1906;Jones,1979, 1981; The taxonomic history of the tribe Vernonieae has Keeley & Jones, 1977). The source of much of this dif- changed over time and the huge or core genus Vernonia ficulty lies with the extensive variability inherent in the Less. was suffering divisions, fragmentations, and rear- group, the occurrence of gradual morphologies and rangements. Based on morphological, phytochemical, overlapping characters with numerous combinations of cytological, palynological, and molecular studies a total autoapomorphic states (Keeley & Jones, 1977;Keeley of 126 genera distributed in 21 subtribes, 15 in the New &Turner,1990; Robinson, 1999). World, and six in the Old World are recognized for the tribe (Keeley & Robinson, 2009). Most of the emerged Correspondence to: Gisela M. Via do Pico. E-mail: gisela_ and resurrected new genera were established based on [email protected] ISSN 1477-2000 print / 1478-0933 online Ó The Trustees of the Natural History Museum, London 2016. All Rights Reserved. http://dx.doi.org/10.1080/14772000.2015.1134699 Floral microcharacters of Chrysolaena (Asteraceae) 225 micromorphological characters. In taxonomic studies on conflictive species of the genus; and (3) determine the sys- American Vernonieae, Robinson (1999) attached great tematic value of these microcharacters in specific segrega- importance to the value of floral microcharacters for sub- tion of the taxa. tribal and generic delimitations (e.g., the style base, the anther appendages, the carpopodium shape, and the shape of cypsela wall crystals). Some of these characters were Materials and methods used as major distinguishing features in the segregation of genera such as Chrysolaena H. Rob., Lessingianthus H. The specimens for this study were obtained from living Rob., Vernonanthura H. Rob., and Lepidaploa H. Rob., material and the herbarium of Instituto de Botanica del previously members of the core genus Vernonia Nordeste (CTES). A complete voucher list is given in the (Robinson, 1988a, 1988b, 1990, 1999). Other characters Appendix. taken into account were the morphology of the pollen For the micromorphological observations with a light grains, chromosome number, chemical composition, and microscope (LM) , florets and fruits were removed from inflorescence shape (Dematteis, 2007, 2009; Robinson, herbarium specimens. Samples were rehydrated in 2009). warm soapy water and dissected under a stereoscopic Chrysolaena belongs to the subtribe Lepidaploinae S. microscope. They were then mounted in Hoyer’s C. Keeley & H. Rob. and comprises 18 species geographi- solution (Anderson, 1954; King & Robinson, 1970). cally concentrated in the central region of South America, Observations were done with a LM equipped with a dig- with the main center of diversity in southern Brazil and ital photographic camera (Canon Power Shot A 640). northeast of Argentina (Dematteis, 2009; Robinson, Trichome types were drawn with a camera lucida 1988b). Most species have a wide geographic distribution, microscope. although there are some endemic entities. The species of Observations with a scanning electron microscope genus are perennial herbs or erect shrubs with well devel- (SEM)were performed on herbarium and living material. oped xylopodia. They are characterized by sericeous or Herbarium samples were rehydrated with warm soapy velutine indumentum, mainly in stems, leaves and phylla- water and fixed with FAA. Disc florets were dehydrated ries, style without basal node, and glandular anthers and in a graded series of acetone and critical point-dried with cypselae (Robinson, 1988b). However these features are liquid CO2. Dried samples were partially dissected, only known for seven species: C. flexuosa (Sims) H. Rob., mounted on stubs, coated with gold/palladium and C. lithospermifolia (Hieron.) H. Rob., C. obovata (Less.) observed with SEM (JEOL 5800 LV). Dematt., C. oligophylla (Vell.) H. Rob., C. platensis For the description of microcharacters, terminology (Spreng.) H. Rob., C. propinqua (Hieron.) H. Rob., and C. proposed by Metcalfe and Chalk (1979), Narayana verbascifolia (Less.) H. Rob. (Robinson, 1988b). Other (1979), Ramayya (1962), and Robinson (1988a, 1988b, distinctive characteristics of the genus are the presence of 1990, 1999) was used. pollen grain type ’C’ (tricolporate, echinolophate with 29 lacunae) and the base chromosome number x D 10 (Dematteis, 2009; Oliveira, Forni-Martins, & Semir, Statistical analyses 2007a, 2007b; Oliveira, Semir, & Forni-Martins, 2012; A data matrix with 59 OTUs £ 13 characters was made. Downloaded by [181.1.176.133] at 07:22 30 January 2016 Via do Pico & Dematteis, 2012, 2013a, 2013b, 2014). The size, the presence of idioblasts and crystals on the Chrysolaena is closely related to Lessingianthus and fruit were not considered because these showed great Lepidaploa, with which it shares some characteristics. intraspecific variation. Also taxa with incomplete infor- With Lepidaploa, for example, it shares the presence of mation were not considered. The characters were coded glands in
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